Argulus Ambystoma, a New Species Parasitic on the Salamander Ambystoma Dumerilii from Mexico (Crustacea: Branchiura: Argulidae)1
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Argulus ambystoma, a New Species Parasitic on the Salamander Ambystoma dumerilii from Mexico (Crustacea: Branchiura: Argulidae)1 WILLIAM J. POLY2, Department of Zoology, Southern Illinois University, Carbondale, IL 62901-6501 ABSTRACT. A new species of Argulus is described based on 18 specimens taken from the salamander ("achoque" or "ajolote") Ambystoma dumerilii Duges, collected in Lake Patzcuaro, Michoacan, Mexico. Diagnostic characters include the shape of the respiratory areas, number of sclerites in suction cup rods, and structures on the legs of males. Females are heavily stippled, whereas males have a very distinctive pigment pattern consisting of abundant melanophores covering the testes dors ally and two dark, inverted triangular patches on the carapace dorsally. The new species is similar to the North American species, A versicolor,A. americanus, A. maculosus, and A diversus. A single, dorsal pore was observed on each caudal ramus using scanning electron microscopy; these pores have not been reported previously in the Branchiura. OHIO J SCI 103 (3>52-6l, 2003 INTRODUCTION 1991; Osorio-Sarabia and others 1986; Perez-Ponce de Ten species of Argulus Miiller have been collected in Leon and others 1994; Peresbarbosa-Rojas and others Mexico, Central America, and the West Indies. The wide- 1994; Espinosa-Huerta and others 1996; Peresbarbosa- spread exotic, Argulus japonicus Thiele, was found on Rojas and others 1997). A number of endemic taxa exist aquarium fishes, Carassius auratus (Linne) and Astro- in Lake Patzcuaro including several fishes (Cbirostoma notus ocellatus (Agassiz), in Puerto Rico (Bunkley-Williams patzcuaro Meek, C. e. estor Jordan, C. a. attenuatum and Williams 1994), and Vargas and Fallas (1976) found Meek), a crayfish (Cambarellus patzcuarensis Villa- A. japonicus on Carassius sp. in Costa Rica in 1972 and lobos), and the salamander host of the new argulid (Am- 1973- Argulus dactylopteri Thorell was described from bystoma dumerilii (Duges)) (Brandon 1970; Barbour specimens collected from the gills of Dactylopterus 1973; Hobbs 1989). volitans (Linne) in the West Indies (Thorell 1865, 1867); neither Cressey (1982) nor Pineda and others (1995) MATERIALS AND METHODS mentioned A. dactylopteri. Wilson (1902) gave the locality Three males and two females were examined using for A. dactylopteri as the East Indian Ocean from the scanning electron microscopy (SEM). Preparation branchial cavity of D. volitans, whereas Yamaguti (1963) procedures were modified slightly from those of Rupp cited Europe and Dactylopterus sp. If the type locality (1990). All specimens were initially preserved and stored of A. dactylopteri is somewhere other than the Atlantic in 70% ethanol (EtOH, J. A. Beatty pers. comm.). Speci- Ocean or Mediterranean Sea, the host would not be D. mens were dehydrated in an EtOH series consisting of volitans but a species of Dactyloptena (see Eschmeyer 80% (5 min), 90% (5 min), 100% (1st, 5 min; 2nd, 10 min), 1997). Only six species of Argulus, A. chromidis Kr0yer; then immediately critical point dried (CO2), mounted A. flavescens Wilson; A. megalops Smith; A. mexicanus on metal stubs with carbon paint, allowed to dry over- Pineda, Paramo, and Del Rio; A. melanostictus Wilson; night in an oven (60° C), and sputter coated with gold/ and A. rhamdiae Wilson have been found in Mexico palladium. Specimens were stored in an oven at 60° C (Wilson 1936; Causey I960; Olson 1972; Cressey 1982; between uses. Some specimens were coated up to three Fucugauchi and others 1988; Pineda and others 1995; times and were examined in a Hitachi S570 Scanning Suarez-Morales and Gasca 1997; Suarez-Morales and Electron Microscope at 15kV. Six females and seven others 1998). In addition, an unidentified species of Argu- males (including the holotype, all adult) were examined lus was reported from Potamarius nelsoni (Evermann under dissecting and compound microscopes in a and Goldsborough) by Pineda-Lopez and others (1985). watchglass or temporary slide mount (with 70% EtOH/ The species described herein was collected from glycerin). The holotype and allotype were photographed Lake Patzcuaro (Lago de Patzcuaro) in Central Mexico, using a camera tube on a light microscope. In addition, State of Michoacan. Both the invertebrate and vertebrate some counts and measurements were taken from 3 male faunas of the lake have been investigated; however, the and 2 female specimens later used for SEM. All genus Argulus has not been recorded as a component measurements were made using an ocular micrometer, of the fauna (Ueno 1939; Ancona and others 1940; and measurements reported in the description are Brehm 1942; Tressler 1954; Brandon 1970; Barbour arranged as follows: range (mean, holotype) with allo- 1973; Rosas and others 1985; Chacon-Torres and others type values substituted in the case of females. Values for the right and left sides refer to the right and left of the specimen in dorsal view. Type specimens were deposited 'Manuscript received 20 September 2001 and in revised form 5 in the National Museum of Natural History, Washington, May 2002 (#01-24). DC (USNM), and specimens of Argulus americanus 2Present Address: Department of Ichthyology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 Wilson (USNM 274331, 274332), A. diversusWihon (USNM OHIO JOURNAL OF SCIENCE W. J. POLY 53 32823,32824), A. maculosusWilsonOJSNM 12226,74329), and A. versicolor Wilson (USNM 74321; MCZ 19604, 19607) were examined by light microscopy for comparative purposes (MCZ, Museum of Comparative Zoology, Harvard University, Cambridge, MA). RESULTS Family Argulidae Rafinesque, 1815 Argulus Miiller, 1785 Argulus ambystoma, new species Material Examined Holotype: adult male, 3-48 mm total length, USNM 282782, Lake Patzcuaro, Michoacan, Mexico. Allotype: adult female, 4.18 mm total length, USNM 282783, col- lected with holotype. Paratypes: 5 adult males, 4 adult females, USNM 282784, collected with holotype; 4 adult males, 3 adult females (used for SEM or cleared, not FIGURE 1. Argulus ambystoma, new species, dorsal view, male (holo- retained), collected with holotype (host collected by type, left) and female (allotype, right) (scale bar = 1.0 mm). J. Arnett; Argulus collected from hosts, Ambystoma dumerilii (Duges), byj. A. Beatty and R. A. Brandon on 27 May 1981). scattered on dorsal surface and margins of carapace. Smaller type of sensillum present on anterior margin Diagnosis (between sinuses) of cephalic region, abundant in inter- Respiratory areas with a smaller, oblong "area" sit- spaces between larger sensilla. Pair of compound eyes uated anterior to and abutting or nearly contacting a anteriorly with diameters (left and right eyes, Jim) 170- larger posterior "area"; respiratory areas outlined with 210 (187, left: 170 and right: 180) (n = 18 eyes) in males, melanophores and very distinct; caudal rami near pos- 200-240 (224, left: 230 and right: 210) (n = 14 eyes) in terior end of abdomen in anal sinus; paired post- females. Transverse distance between eyes (|Xm) 390- antennal spines with reduced anterior spine and stout 440 (419, 410) in males, 510-650 (604, 570) in females. posterior spine; males with 43-56 support rods per suc- Nauplius eye with one anterior and two posterior ocelli, tion cup; females with 45-54 support rods per suction pigment between ocelli forming a dark, prominent "Y" cup; usually 3 to 4 sclerites per rod in males and 4 to 5 or "V." Sclerotized dorsal ridges forked anterior of eyes, sclerites per rod in females; basal plate of second maxilla inner branches longer and less developed than outer with three stout, digitate spines; first two pairs of legs branches. Bifurcation of dorsal ridges usually more ob- each with a recurved flagellum on dorsal surface; coxae vious in males and sometimes appears absent in females. of second legs of males with bilobate structure pos- Ventrally, carapace with small, posteriorly-projecting teroventrally, scales on lobes distally; third legs of males spines along outer margin, more numerous anterior of with paddle-like extension issuing anterodistally from respiratory areas with a few rows extending along outer the coxae, extending over bases. margin of carapace and respiratory areas to level of second legs. Respiratory areas consist of smaller, oblong Description "area" situated anterior to and abutting or nearly con- Total length (mm) 3.26-3.72 (3.53, 3.48) in males, tacting larger posterior "area"; respiratory areas distinct, 4.06-5.85 (4.92, 4.18) in females. Carapace ovoid to outlined with melanophores (Fig. 3). circular, narrower anteriorly. Carapace length (mean of Thorax dorsoventrally compressed, indistinctly seg- both sides, mm) 2.16-2.47 (2.33, 2.26) in males, 3.20- mented with two pairs of posteriorly-projecting spines 3-97 (3.57, 3-22) in females. Maximum carapace width ventrally. Spines digitate, rounded distally, the anterior (mm) 2.13-2.57 (2.41, 2.38) in males, 3.21-4.15 (3-66, pair (accessory spines) larger than posterior pair 3-30) in females. Carapace extending to middle of third (postmaxillary spines). Accessory spines situated be- legs or posterior edge of third legs/anterior margin of tween basal segments of second maxillae; postmaxillary abdomen in males. Carapace extending to middle of spines positioned farther apart than accessory spines. third legs or posterior of fourth legs/anterior of abdo- Thorax with coarse-pectinate scales scattered on ventral men in females, varies considerably in females. Females surface. Four pairs of biramous swimming